Effects of the toxic dinoflagellate <Emphasis Type="Italic">Gymnodinium catenatum</Emphasis> on uptake and fate of paralytic shellfish poisons in the Pacific giant lions-paw scallop <Emphasis Type="Italic">Nodipecten subnodosus</Emphasis>

Juvenile Pacific giant lions-paw scallops Nodipecten subnodosus were fed the toxic dinoflagellate Gymnodinium catenatum, a producer of paralytic shellfish poison (PSP), supplied with Isochrysis galbana (a nontoxic microalgae). Short-term (<24 h) experiments were performed to determine clearance and ingestion rates of G. catenatum. Kinetics of PSP was examined in longer-term experiments (>2 days). At high food concentrations, juvenile scallops showed production of pseudofeces, partial shell valve closure, and reduction in feeding. According to HPLC analysis, the only toxins present in the dinoflagellate G. catenatum and in the scallops were the gonyautoxins (GTXs), except in the labial palps and digestive gland, where trace amounts of saxitoxin (STX) were present in scallops. These tissues could play an important role in toxin biotransformation. The ranking of toxin concentration in tissues was: digestive gland > labial palps > intestine > gills > mantle > adductor muscle, where the total contribution of viscera was more than 80% of the total toxin body burden. Juvenile scallops exhibited no apparent detrimental physiological responses during the long-term feeding experiment. The dinoflagellate may contribute nutrients to the scallop, in addition to the microalgae I. galbana. The dinoflagellate may enhance cell uptake and byssus production. Once PSP accumulated during the first 12 days, it was slowly eliminated. The limited capacity for accumulating toxins in the adductor muscle favors domestic marketing of scallops.     

Field depuration and biotransformation of paralytic shellfish toxins in scallop<Emphasis Type="Italic"> Chlamys nobilis</Emphasis> and green-lipped mussel<Emphasis Type="Italic"> Perna viridis</Emphasis>

Under laboratory conditions, the scallop Chlamys nobilis and the mussel Perna viridis were exposed to N-sulfocarbamoyl toxins (C2 toxin), a paralytic shellfish toxin (PST), by feeding a local toxic strain of the dinoflagellate Alexandrium tamarense (ATDP) that produced C2 toxin exclusively. The bivalves were subsequently depurated in the field, and their depuration kinetics, biotransformation and toxin distribution were quantified. Depuration was characterized by a rapid loss within the first day, followed by a secondary slower loss of toxins. In the fast depuration phase, scallops detoxified PSTs more quickly than the mussels (depuration rate constants for scallops and mussels were 1.16 day^–1 and 0.87 day^–1, respectively). In contrast, the mussels detoxified PSTs more quickly than the scallops in the slow depuration phase, and the calculated depuration rate constants (mean+SE) from day 2 to day 13 were 0.063+0.009 day^–1 and 0.040+0.019 day^–1 for mussels and scallops, respectively. The differences in the appearances of gonyautoxins, GTX2 and GTX3, and their decarbamoyl derivatives, dcGTX2, dcGTX3 and GTX5, which are all derivatives of C2 toxin, indicated active and species-specific biotransformation of the algal toxins in the two bivalves. In both species of bivalves, the non-viscera tissue contained fewer toxins and lower concentrations than the viscera-containing tissue compartment. In scallops, very little toxin was distributed in the adductor muscle. In mussels, most of the PSTs were found in the digestive gland with significant transport of toxins into the digestive gland from other tissues during the course of depuration. The toxin profiles of scallops and mussels differed from each other and from that of the toxic algae fed. A significant fraction of GTX5 was detected in the mussels but not in the scallops. Our study demonstrates a species specificity in the depuration kinetics, biotransformation and tissue distribution of PSTs among different bivalves.Communicated by T. Ikeda, Hakodate     

Toxin accumulation and feeding behaviour of the planktonic copepod Calanus finmarchicus exposed to the red-tide dinoflagellate Alexandrium excavatum

The planktonic copepod Calanus finmarchicus is a dominant member of the zooplankton community in the lower St. Lawrence Estuary in eastern Canada. Blooms of the toxic marine dinoflagellate Alexandrium excavatum which produces high cellular levels of paralytic shellfish poisoning (PSP) toxins, occur during the period of high C. finmarchicus production in summer in this region. To study the feeding behaviour of C. finmarchicus in the presence of Alexandrium spp., experiments were conducted in which female adult copepods collected from the St. Lawrence Estuary between May and September 1991 were exposed under controlled conditions to two toxic isolates of A. excavatum (Pr18b and Pr11f) from the estuary and to a non-toxic control (PLY 173) of a closely related species, A. tamarense isolated from the Tamar Estuary, Plymouth, U.K. Clearance rates on non-toxic A. tamarense cells averaged 5.5 ml ind^-1 h^-1 but were nearzero with either toxic isolate. When presented with a mixture of A. excavatum and the non-toxic diatom Thalassiosira weissflogii in varying proportions, C. finmarchicus fed upon the diatom but avoided the toxic dinoflagellate. Although feeding rates on A. excavatum were very low, toxin analysis by high-performance liquid chromatography with fluorescence detection (HPLC-FD) revealed that the PSP toxins were accumulated in copepods exposed to toxigenic dinoflagellates.The toxin composition in copepods was similar to that of the toxic dinoflagellate, but not necessarily identical, particularly after short-term (2-h) exposure, when relatively elevated levels of N-sulfocarbamoyl toxins were detected. The evidence suggests that C. finmarchicus ingests toxic dinoflagellate cells, either mistakenly or during exploratory bouts of feeding, and accumulates PSP toxins in its gut system and perhaps in other tissues.     

Deleterious effects of a nonPST bioactive compound(s) from <Emphasis Type="Italic">Alexandrium tamarense</Emphasis> on bivalve hemocytes

The known negative effects of shellfish toxin-producing dinoflagellates on feeding, burrowing and survival of some bivalve mollusks has prompted questions concerning whether they might also impair the internal defense system of affected bivalves and make them more susceptible to disease agents. The primary components of the cellular defense system are hemocytes. Many toxic dinoflagellates are too large to be ingested whole by hemocytes and would most likely be exposed to intracellular toxins only after the algae are consumed, broken down, and the water-soluble toxins, released. Therefore, we conducted a series of experiments in which hemocytes from two suspension-feeding bivalves—the Manila clam, Ruditapes philippinarum, and the softshell clam, Mya arenaria—were exposed in vitro to filtered extracts of one highly toxic paralytic shellfish toxin (PST)-producing and one nonPST-producing strain of Alexandrium tamarense (isolates PR18b, 76 ± 6 STXeq cell⁻¹ and CCMP115, with undetectable PST, respectively). We measured adherence and phagocytosis, two hemocyte attributes known to be inhibited by bacterial pathogens and other stressors. We found no measurable effect of a cell-free extract from a highly concentrated suspension of the PST-producing strain on hemocytes of either bivalve species. Instead, extract from the nonPST-producing strain had a consistent negative effect on both clams, resulting in significantly lower adherence and phagocytosis compared to strain PR18b and filtered seawater controls. The bioactive compound produced by strain CCMP115, which has yet to be characterized, may be similar to the PST-independent allelopathic compounds described for Alexandrium spp., which act on other plankters. These compounds and those produced by other harmful algae are known to cause immobilization, cellular deformation and lysis of co-occurring target organisms. Thus, nonPST producing Alexandrium spp., which do not cause paralysis and burrowing incapacitation of clams, may still produce a compound(s) that has negative effects not only on hemocytes, but on other molluscan cell types and their functions, as well.     

Accumulation of paralytic shellfish poisoning toxins in planktonic copepods during a bloom of the toxic dinoflagellate<Emphasis Type="Italic"> Alexandrium tamarense</Emphasis> in Hiroshima Bay,western Japan

Concentrations of paralytic shellfish poisoning (PSP) toxins in toxic dinoflagellate cells and in marine planktonic copepods were monitored during the bloom of Alexandrium tamarense in Hiroshima Bay, western Japan. Concentration of the toxins retained by copepods was a function of the ambient toxin concentration, i.e. the product of A. tamarense cell density and cellular toxicity. The toxin concentration in copepods increased with the increase of toxicants in the seawater then leveled off, but decreased significantly at higher concentrations. In the field, the maximum toxin concentration was 1.2 pmol ind^-1, whereas in the laboratory, the copepod Acartia omorii accumulated a much higher concentration of PSP toxins (24 pmol ind^-1). Feeding avoidance against Alexandrium tamarense and a shift to alternative food sources such as diatoms in the field might keep their toxin levels lower than their potentially maximum level. The copepod toxin levels in the field were not so high as to cause an instantaneous lethal effect on their predator fishes but may reach possibly lethal levels after a few days' continuous feeding. Overall toxin retention by copepods after 12 h feeding and 2 h starvation was only 2.5% of total ingested toxins, which suggested that a significant amount of toxins was released into the seawater. Measurements of toxin reduction and gut evacuation suggested that the toxins were removed through both fecal evacuation and metabolism (e.g. excretion, decomposition and transformation). The results, as a whole, imply that copepods can be a link for PSP toxin flux in both pelagic and benthic food webs and can also be a sink for toxins by metabolizing and removing them from the environment.Communicated by T. Ikeda, Hakodate     

Effects of induced paralysis on hemocytes and tissues of the giant lions-paw scallop by paralyzing shellfish poison

In general, bivalves are not affected by exposure to toxic dinoflagellates that produce paralyzing shellfish poisons (PSP). After injection with PSP extracted from the Gymnodinium catenatum, Nodipecten subnodosus is paralyzed, indicating that PSP provokes effects similar to what is observed in vertebrates, including paralysis and metabolic stress. To investigate the processes involved in poisoning by PSP, lions-paw scallops were injected with gonyautoxin (GTX) 2/3 epimers in the adductor muscle. Mild doses provoked adductor muscle contractions and paralysis, mantle retraction, and incapacity of shell closure, but scallops gradually recovered in a clear, dose-time recovery pattern. With high doses of GTX 2/3, scallops were permanently paralyzed, and hemocytes in hemolymph were reduced. Surprisingly, under these conditions, scallops continued normal feeding and did not show any microscopic defect in intestine or gills, but hemocytes infiltrated the adductor muscle and abnormal vitellogenesis and mantle melanization occurred. Paralysis stress was accompanied by negative scallop responses, based on visible effects, generation of nitric oxide, lipid peroxidation, and changes in antioxidant and hydrolytic enzymes in hemocytes and tissues. These data can be used to understand potential side effects of PSP in bivalves.     

Postingestive selection in the sea scallop (Placopecten magellanicus) on the basis of chemical properties of particles

Suspension-feeding bivalves are adapted to use a highly variable mixture of particles by sorting them before and after ingestion. Postingestive sorting in bivalves has been confirmed for several bivalve species, but few studies have attempted to isolate the factors influencing postingestive selection among different particles presented simultaneously. The ability of the sea scallop, Placopecten magellanicus (Gmelin), to sort a mixture of organic (¹⁴C-labeled dinoflagellates, Prorocentrum minimum) and inorganic (⁵¹Cr-labeled beads, diameter 16-18 µm) particles of similar size and shape within the stomach was examined. The study was carried out in August 1997 using scallops collected near Deer Island, New Brunswick, Canada. Sorting of the two particle types was measured by dissecting the scallops at intervals after feeding and comparing the ¹⁴C:⁵¹Cr ratios within the stomach, digestive gland, and feces. The ¹⁴C:⁵¹Cr ratio in the stomach decreased over time, indicating that sea scallops were sorting organic from inorganic particles. The ability of P. magellanicus to sort particles solely on the basis of chemical properties was tested in a second study by presenting them with a mixture of protein-coated and uncoated beads of two different colors. This study was carried out in February 2000 using scallops collected near Maces Bay, New Brunswick, Canada. Gut retention times of the two types of beads were measured using flow cytometry. Scallops retained protein-coated beads in the gut longer than uncoated beads, indicating postingestive selection by chemical properties. This study provides the first evidence for simultaneous postingestive sorting by bivalves of particles based solely on chemical properties. This would potentially enable P. magellanicus to preferentially retain particles of higher food quality longer than those of poor quality, thereby enhancing digestive efficiency.     

镉胁迫对不同镉富集能力海水养殖贝类抗氧化能力的影响——以扇贝和菲律宾蛤仔为例

为研究具有不同镉(Cd)富集能力的扇贝和菲律宾蛤仔在相同镉胁迫环境下的抗氧化能力反应的差异,将2种贝类于0.05mg·L-1Cd环境中暴露10 d,分别于第0天、2天、4天、6天、8天、10天分别取样检测内脏团中SOD、CAT、GPx、GST酶活性和GSH含量,同时分析内脏团中Cd的含量。结果表明,对于Cd胁迫组,扇贝和菲律宾蛤仔内脏团中SOD和CAT酶活反应相似,均呈先被诱导后被抑制的规律,且2种贝类的SOD和CAT活性差异较小。2种贝类内脏团中GSH含量、GST和GPx活性变化差异较大:扇贝内脏团中GSH含量显著降低(p0.05),GST和GPx活性均在第2天和第4天时处于显著诱导状态(p0.05),从第6天时处于抑制状态,而菲律宾蛤仔内脏团中GSH含量、GST和GPx活性在Cd暴露期间无显著变化,且2种贝类间比较,扇贝内脏团中GSH含量和GST活性在整个实验期间始终显著高于菲律宾蛤仔(p0.05),GPx活性从第4天开始高于菲律宾蛤仔;对于无镉污染的对照组,扇贝和菲律宾蛤仔内脏团中SOD、CAT和GPx活性无显著差异,但扇贝内脏团中GSH含量和GST活性显著高于菲律宾蛤仔,其中前者GSH含量约为后者的22倍。研究同时表明在相同镉环境下扇贝内脏团对Cd的富集浓度和富集速率均远高于菲律宾蛤仔。本研究明确了2种贝类在相同镉环境下的抗氧化反应的差异,推断出扇贝内脏团中高含量的GSH以及较高的GST和GPx活性可能在扇贝高富集、高耐受Cd方面起重要作用。     

Comparison of feeding and biodeposition of three bivalves at different food levels

Experiments on the edible mussel Mytilus edulis, the American oyster Crassostrea virginica, and the hard clam Mercenaria mercenaria, using flowing systems, showed that the feeding and biodeposition rates were affected by food concentration. At all levels of food concentration, the order of increasing feeding rate (both the percent of available particulate carbon and the actual amount of carbon removed) was: clam < oyster < mussel. All bivalves exhibited lower feeding rates (both precent and actual) at low food concentrations. However, the precent of available food removed quickly increased to a maximum at food concentrations typical for the natural environment. This maximum remained constant for the mussel and oyster, but declined with increasing food concentration for the clam. However, because this percentage was for increasing levels, the actual carbon removed continued to increase up to the highest food level for all three bivalves. In increasing order of biodeposition rate, the bivalves were: clam < oyster < mussel. The biodeposition rates of the three bivalves increased logarithmically with increased food concentration as a result of the production of pseudofeces. The feeding and biodeposition data were used to calculate assimilation rates, and this pointed out the higher efficiency of the oyster compared to the mussel and clam.Contribution No. 2993 from the Woods Hole Oceanographic Institution.     

Ontogenetic changes in gill morphology and potential significance for food acquisition in the scallop <Emphasis Type="Italic">Placopecten</Emphasis><Emphasis Type="Italic">magellanicus</Emphasis>

Sources of mortality in both wild and cultured populations of marine bivalves during postlarval stages remain largely unknown, but may be partly associated with the inability to meet energetic demands during intense morphogenesis. The development of the gills in postsettlement scallops (Placopecten magellanicus) from 0.35 to 14 mm in shell height (SH) was investigated using scanning electron microscopy to determine the degree of size-specific differentiation of the gills and evaluate potential ontogenetic constraints in food acquisition. Key transitional stages in morphogenesis, likely to exert pronounced effects on feeding function, were identified and correlated with scallop size. The gill was initially homorhabdic, with unreflected inner demibranchs forming a basket-like structure maintained by ciliary junctions. Gill reflection, immediately followed by accelerated proliferation of gill filaments and formation of outer demibranchs, occurred at ~1 mm SH. Outer demibranchs were fully formed at ~2 mm SH. Suspension-feeding is probably rather inefficient prior to attaining 1-2 mm sizes. The onset of the heterorhabdic, adult form of the gill, which allows bidirectional particle transport and the potential for selection and for volume regulation of ingested material on the gill, occurred fairly late in development, at ~3.3-5.0 mm SH. Full development of gill plication was delayed until scallops attained ~7 mm. Gill differentiation in this species is thus relatively protracted and punctuated by critical transitional stages, which may be important in determining feeding and growth capacity of postlarval wild and cultured populations.     

Trace metal ingestion and assimilation by the green mussel <Emphasis Type="Italic">Perna viridis </Emphasis>in a phytoplankton and sediment mixture

We examined trace metal ingestion and assimilation (Cd, Se, Zn) by the green mussel Perna viridis in a mixture of diatoms and sediment at concentrations below and above the pseudofeces production levels. Dual gamma radiotracers (¹⁰⁹Cd, ⁶⁵Zn) were used to investigate particle selection on marine diatoms and sediments. The diatom (Thalassiosira weissflogii) was radiolabeled with ¹⁰⁹Cd, and the natural sediment was radiolabeled with ⁶⁵Zn. By comparing the ratios of ¹⁰⁹Cd:⁶⁵Zn in the particle mixture, mussel tissues and pseudofeces within a short-term exposure period (35 min), the results demonstrated that the green mussels were able to selectively ingest the diatom particles at a high particle load. Efficiency of selection for nutritious particles (e.g. diatoms) increased with increasing ratio of sediments in the particle mixture. Pseudofeces contained a higher ratio of sediments relative to that in the feeding suspension. No major particle selection was observed at concentrations below the level for pseudofeces production. The assimilation of Cd, Se and Zn by the green mussels was quantified using a pulse-chase feeding technique. The assimilation of Se and Zn by the green mussels from ingested diatoms was reduced with the presence of sediment within the mussel gut, presumably due to the resorption of metals onto the sediment, leading to a quicker passage of metals through the digestive tract and a lower proportion of metals subjected to intensive digestion. In contrast, the presence of diatoms did not significantly affect metal assimilation from ingested sediment. A significant correlation between metal assimilation efficiency and metal gut passage time was also observed. Metal assimilation by the green mussels appeared to be little dependent on the particle concentration in seawater. Our study suggests that particle selection may potentially alter metal influx from ingested food sources, particularly at high particle concentrations. Selective feeding of nutritious particles, coupled with a high assimilation efficiency from these ingested particles, may increase metal influx into mussels from the dietary phase.     

Observations on shell deformities,ultrastructure, and increment formation in the bay scallop Argopecten irradians

Shell morphology and ultrastructure were examined in the bay scallop Argopecten irradians, cultured in recirculating seawater systems under various conditions of feeding, lighting, and handling. On a unialgal diet of Thalassiosira pseudonana, scallop growth ranged from 120 to 183 m d^-1 at 20°C in the laboratory, about two-thirds of the growth rate found in the field. However, shell deposited in the laboratory differed from that in the field in several ways. In the field, scallops formed costae as an unpigmented, corrugated marginal shell layer; shell deposited in the laboratory lacked this layer and was therefore darker. Also, microstructure of the exterior shell surface of field scallops was coarsely granular, while that of cultured scallops was relatively smooth. Excessive handling of scallops in the laboratory resulted in marginal thickening of valves, a deformity which was completely arrested by a change from daily to weekly handling. Scallops cultured in the same tank with oysters developed shell-thickening on the interior of the valves. It is postulated that shell abnormalities in bivalves result from disruption of complex behavioral processes associated with shell deposition and may be elicited by a variety of natural and experimental irritants. Under natural lighting regimes and optimal conditions for growth, scallops deposited exactly one shell increment per day, but under continuous lighting, deposition of growth increments often became aphasic. In one 28-d experiment, there was a strong correlation between number of growth increments formed and increase in shell height, suggesting that shell ridge formation occurred intermittently, rather than daily, when shell growth rates fell below approximately 150 m d^-1.     

Feeding responses of the bivalves Crassostrea gigas and Mytilus trossulus to chemical composition of fresh and aged kelp detritus

The chemical composition of kelps (e.g. polyphenolics) deters grazing by herbivores, but kelp detritus is potentially a source of nutrition for suspension feeders. The effects of kelp detritus derived from two species [Agarum fimbriatum Harvey and Costaria costata (Turner) Saunders] on feeding of oysters, Crassostrea gigas Thunberg, and mussels, Mytilus trossulus Gould, were examined in feeding experiments. Fresh and aged kelp particles were sequentially presented in combination with the microalga Rhodomonas lens at an initial total concentration of 5᎒^-4 ml^-1. Aging of kelp particles for 4 days in seawater significantly reduced the concentration of polyphenolics without changing the total carbon or nitrogen content. Clearance rates of both mussels and oysters were significantly lower in the presence of fresh versus aged kelp particles, and clearance rates declined overall with declining polyphenolic concentrations. Video endoscopy was used to examine feeding selectivity at the level of the gill in oysters in the same food treatments used in the clearance rate experiments. Comparison of particle composition in the water versus the pseudofeces in both oysters and mussels was also used as a measure of feeding selectivity. When presented with R. lens in combination with fresh and aged kelp particles selectivity for R. lens tended to be greater against fresh than aged particles, and there was some indication that this was stronger for A. fimbriatum than for C. costata particles. The ability to select was lower at very high polyphenolic concentrations, which may reflect poisoning of sensory binding sites. These data suggest that bivalves distinguish among particles of varying chemical composition and respond by changing their clearance rates and their selectivity.     

Paralytic shellfish poison in the “hiogi” scallop Chlamys nobilis

Attempts were made to analyze the toxin composition of the toxic hiogi scallop Chlamys nobilis. The toxins were partially purified from the digestive glands by column chromatography using Bio-Gel P-2 and Bio-Rex 70 (H⁺ form), resulting in separation into protogonyautoxin (PX), gonyautoxin (GTX) and saxitoxin (STX) fractions. Their total potencies were scored to be <100 mouse units (MU), 3200 MU and 3700 MU, respectively. A 5-min hydrolysis with 0.1 N HCl enhanced the potencies of PX and GTX fractions to 450 MU and 12000 MU respectively, whereas no enhancement occurred in the STX fraction at all. Electrophoretic, thin-layer chromatographic and high performance liquid chromatographic analyses demonstrated that the PX fraction consisted mainly of GTX₈ and its epimer, the GTX fraction of GTX₅, GTX₆, along with two unidentified toxins, and the STX fraction exclusively of two unidentified toxins. This rather unique composition suggested a complex metabolism of PSP in this species.     

Mortality and growth of juvenile hard clams Mercenaria mercenaria during brown tide

Blooms of the picoplankton-sized alga Aureococcus anophagefferens Hargraves et Sieburth, also known as "brown tide," frequently recur in Great South Bay, New York, USA. A field study compared mortality and growth rates of juvenile hard clams, Mercenaria mercenaria (L.), raised at a site that experienced a bloom of A. anophagefferens (>1᎒⁶ cells ml^-1) to clams grown in a nonbloom site from May to September 2000 at two sites on the north and south shores of Long Island. High (67%) cumulative clam mortality was observed for clams raised at the brown tide site (south shore) during the bloom (June-July), and individuals exhibited no measurable growth during that period. After the bloom subsided (mid-July), clams that survived the brown tide suffered low (<5%) cumulative mortality over the remaining sampling period (July-September). These survivors exhibited rapid growth for 4 weeks shortly after the bloom's decline. Subsequently, their growth slowed to rates comparable with those observed at the nonbloom site (north shore). By contrast, a low (<5%) cumulative mortality was observed for M. mercenaria raised at the site with no brown tide, and positive growth occurred throughout the study. These data suggest that some juvenile clams not only survive brown tide but also recover following the bloom.     

Discard analysis and damage to non-target species in the "rapido" trawl fishery

This paper describes the catch composition in the rapido trawl fishery and the direct effects on non-target species. All data were collected on commercial fishing vessels so as to reflect commercial rapido-trawling practice. The effects on non-target species were measured using two different damage scales (three- and seven-level scales) depending on the morphology of the taxa. Damage assessment was performed taking into account the whole fishing process by collecting individuals that passed through the cod-end, individuals that were retained in the cod-end and dropped onto the deck and individuals that were collected at the end of the sorting operation just before their return to the sea. Due to differences in the habitat and spatial distribution of target species, discard/commercial ratio was very different among the three different target species fisheries: 1:6 in the queen scallop (Aequipecten opercularis) fishery, 2:1 in the flatfish (Solea spp., Platichthys flesus, Psetta maximus and Scophthalmus rhombus) fishery and 9:1 in the scallop (Pecten jacobaeus) fishery. Damage sustained by non-target species was species-specific and related to the morphology of different organisms. The sorting operation produced similar levels of injury to those of the gear itself: all discarded animals showed higher levels of damage after the sorting than before. Damage to animals that had passed through the cod-end followed the same pattern, and these data could give an estimate of the "unobserved mortality". Our observations indicated a higher impact on non-target species caused by the queen scallop fishery than that caused by the flatfish fishery. This could be due to the total amount of hard-shelled species (in the queen scallop fishery, A. opercularis accounted for 87% of the total catch biomass) in any given haul, since shells macerated the catch during towing. Discarded animals from the queen scallop fishery showed higher levels of damage than those collected in the flatfish fishery. The rapido trawl fishery seemed to exert a strong selective pressure on the macrobenthic community, being able to modify the epibenthic fauna structure which, in heavily exploited fishing grounds, was dominated by bivalves, gastropods, crabs, starfish and brittlestars.     

Transfer of brevetoxins to a tellinid bivalve by suspension- and deposit-feeding and its implications for clay mitigation of <Emphasis Type="Italic">Karenia brevis</Emphasis> blooms

Blooms of the brevetoxin-producing Karenia brevis in the Gulf of Mexico cause massive fish kills, food poisoning and adverse respiratory effects in humans. Sedimentation of toxic cells following inert clay application could reduce toxin incorporation by commercially important suspension-feeding bivalves and thus prevent direct public health impacts, but could potentially lead to brevetoxin (PbTx) accumulation by benthic deposit-feeders. The goal of this study was therefore to compare suspension- and deposit-feeding as pathways for brevetoxins. We investigated: (1) the effect of toxic K. brevis on both feeding modes using a facultative deposit-suspension feeding tellinid bivalve, the clam Macoma balthica, as a model species and (2) the relative effectiveness of brevetoxin transfer via suspension- and deposit-feeding over 24-h exposure. Sedimentation of K. brevis was achieved by treatment with 0.25 g phosphatic clay l⁻¹ and brevetoxin concentrations were measured by ELISA. Karenia brevis reduced both suspension- and deposit-feeding activity. This study demonstrates that brevetoxins can be rapidly accumulated by a surface deposit-feeding bivalve from sedimented K. brevis cells and that comparable toxin levels can be attained by both suspension- and deposit-feeding modes [1.2–1.6 μg PbTx (g tissue wet weight)⁻¹]. Deposit-feeding clams generally do not pose a direct threat to humans but may provide a pathway for brevetoxin food web transfer.     

The role of marine aggregates in the ingestion of picoplankton-size particles by suspension-feeding molluscs

Suspension-feeding molluscs are important members of coastal communities and a large body of literature focuses on their feeding processes, including the efficiency of particle capture. Some molluscs, such as bivalves, capture individual picoplankton cells (0.2–2.0 μm) with a retention efficiency of less than 50%, leading to the assumption that such particles are not an important food resource. Picoplankton, however, are often concentrated in particle aggregates of much larger size. This study investigates the ability of suspension feeders to ingest picoplankton-size particles (0.2–2.0 μm) bound in marine aggregates. We fed clams (Mercenaria mercenaria), mussels (Mytilus edulis), oysters (Crassostrea virginica), scallops (Argopecten irradians) and slipper snails (Crepidula fornicata) 1.0- and 0.5-μm fluorescent particles (either polystyrene beads or bacteria) that were (1) dispersed in seawater, or (2) embedded within laboratory-made aggregates. Dispersed 10-μm beads were also delivered so that feeding activity could be determined. Ingested fluorescent particles were recovered in feces or isolated digestive glands and quantified. Results indicate that aggregates significantly enhance the ingestion of 1.0- and 0.5-μm beads by all species of bivalves, and enhance the ingestion of bacteria (greatest cell dimension ca. 0.6 μm) by all suspension feeders examined. Differences among species in their ability to ingest aggregates and picoplankton-size particles, however, were evident. Compared to mussels and clams, scallops and oysters ingested fewer aggregates with 1.0-μm beads or bacteria, and slipper snails ingested the most dispersed beads and bacteria. These differences may be a consequence of variations in gill structure and mechanisms of particle processing. Our data demonstrate that suspension feeders can ingest picoplankton-size particles that are embedded within aggregates, and suggest that such constituent particles may be an important food resource.     

The behaviour of giant clams (Bivalvia: Cardiidae: Tridacninae)

Giant clams, the largest living bivalves, live in close association with coral reefs throughout the Indo-Pacific. These iconic invertebrates perform numerous important ecological roles as well as serve as flagship species—drawing attention to the ongoing destruction of coral reefs and their associated biodiversity. To date, no review of giant clams has focussed on their behaviour, yet this component of their autecology is critical to their life history and hence conservation. Almost 100 articles published between 1865 and 2014 include behavioural observations, and these have been collated and synthesised into five sections: spawning, locomotion, feeding, anti-predation, and stress responses. Even though the exact cues for spawning in the wild have yet to be elucidated, giant clams appear to display diel and lunar periodicities in reproduction, and for some species, peak breeding seasons have been established. Perhaps surprisingly, giant clams have considerable mobility, ranging from swimming and gliding as larvae to crawling in juveniles and adults. Chemotaxis and geotaxis have been established, but giant clams are not phototactic. At least one species exhibits clumping behaviour, which may enhance physical stabilisation, facilitate reproduction, or provide protection from predators. Giant clams undergo several shifts in their mode of acquiring nutrition; starting with a lecithotrophic and planktotrophic diet as larvae, switching to pedal feeding after metamorphosis followed by the transition to a dual mode of filter feeding and phototrophy once symbiosis with zooxanthellae (Symbiodinium spp.) is established. Because of their shell weight and/or byssal attachment, adult giant clams are unable to escape rapidly from threats using locomotion. Instead, they exhibit a suite of visually mediated anti-predation behaviours that include sudden contraction of the mantle, valve adduction, and squirting of water. Knowledge on the behaviour of giant clams will benefit conservation and restocking efforts and help fine-tune mariculture techniques. Understanding the repertoire of giant clam behaviours will also facilitate the prediction of threshold levels for sustainable exploitation as well as recovery rates of depleted clam populations.     

Interactions between two introduced species: Zostera japonica (dwarf eelgrass) facilitates itself and reduces condition of Ruditapes philippinarum (Manila clam) on intertidal flats

Dwarf eelgrass (duckgrass; Zostera japonica) and Manila clams (Ruditapes philippinarum) are two introduced species that co-occur on intertidal flats of the northeast Pacific. Through factorial manipulation of clam (0, 62.5, 125 clams m⁻²) and eelgrass density (present, removed by hand, harrowed), we examined intra- and interspecific effects on performance, as well as modification of the physical environment. The presence of eelgrass reduced water flow by up to 40% and was also observed to retain water at low tide, which may ameliorate desiccation and explain why eelgrass grew faster in the presence of conspecifics (positive feedback). Although shell growth of small (20–50 mm) clams was not consistently affected by either treatment in this 2-month experiment, clam condition improved when eelgrass was removed. Reciprocally, clams at aquaculture densities had no effect on eelgrass growth, clam growth and condition, or porewater nutrients. Overall, only Z. japonica demonstrated strong population-level interactions. Interspecific results support an emerging paradigm that invasive marine ecosystem engineers often negatively affect infauna. Positive feedbacks for Z. japonica may characterize its intraspecific effects particularly at the stressful intertidal elevation of this study (+1 m above mean lower low water).